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Pleistocene Gobiid Fishes of the Genus Rhinogobius

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Bull. Kitakyushu Mus. Nat. Hist., 7: 111-119. December 20, 1987<br />

<strong>Pleistocene</strong> <strong>Gobiid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>the</strong> <strong>Genus</strong> <strong>Rhinogobius</strong><br />

from Kusu Basin, Oita Prefecture, Japan<br />

Yoshitaka Yabumoto<br />

Kitakyushu Museum <strong>of</strong> Natural History, Nishihonmachi,<br />

Kitakyushu, 805 Japan<br />

(Received August 21, 1987)<br />

Abstract Two species <strong>of</strong> <strong>Pleistocene</strong> gobiid fishes, <strong>Rhinogobius</strong> giurinus and R. brumous,<br />

were collected from <strong>the</strong> diatomite bed at Kusu Basin in Oita Prefecture, Japan. These<br />

species are identified by characters <strong>of</strong> <strong>the</strong> pelvic fin and scales. From Kusu Basin, a<br />

salmonid species, Oncorhyncus cf. 0. masou or 0. rhodums, and three cyprinid species<br />

Hemibarbus barbus, Zacco temminckii, and Acheilognathus lanceolata were reported by Uyeno et<br />

al. (1975). This is <strong>the</strong> first record <strong>of</strong> gobiid fishes which were found from this deposit.<br />

A number <strong>of</strong> fossil specimens belonging to <strong>the</strong> family <strong>Gobiid</strong>ae were collected from<br />

Nogami Member in Kusu Formation at Nogami in Kusu Basin, Oita Prefecture. This<br />

Formation and Member were first described by Shuto (1953) who made a geological<br />

survey <strong>of</strong> <strong>the</strong> upper area <strong>of</strong> Kusu River, and named it as Kusu Formation and recognized<br />

three Members: Hosenji, Nogami, and Hijiu Member in order from bottom to top. The<br />

age <strong>of</strong> Nogami Member is Middle <strong>Pleistocene</strong> (Iwauchi and Hase, 1987). Uyeno, et al.<br />

(1975) reported freshwater fish fossils from Nogami Member. These are a salmonid<br />

species, Oncorhyncus cf. 0. masou or 0. rhodums, and three cyprinid species, Hemibarbus<br />

barbus, Zacco temminckii, and Acheilognathus lanceolata.<br />

Two species <strong>of</strong> gobiid fossil fishes <strong>of</strong><br />

<strong>the</strong> genus <strong>Rhinogobius</strong> are reported here for <strong>the</strong> first time from Kusu Basin.<br />

Acknowledgments<br />

I am very grateful to Dr. Teruya Uyeno <strong>of</strong> <strong>the</strong> National Science Museum for his<br />

invaluable advice and critical reading <strong>of</strong> <strong>the</strong> manuscript. I would like to express my<br />

sincere gratitude to Mr. Akinori Takayama <strong>of</strong> <strong>the</strong> Hakusan Kogyo Corporation for <strong>the</strong><br />

donation <strong>of</strong> his specimens and his cooperation in collecting o<strong>the</strong>r specimens. I thank Mr.<br />

Tsugito Nagashima for <strong>the</strong> first information <strong>of</strong> <strong>the</strong> yielding <strong>of</strong> <strong>the</strong> specimens. I am<br />

grateful to Dr. Nobuaki Mizuno <strong>of</strong> Ehime University for <strong>the</strong> donation <strong>of</strong> some forms <strong>of</strong><br />

Recent specimens <strong>of</strong> R. brunneus, and to Mr. Muneyoshi Hayashi <strong>of</strong> <strong>the</strong> Yokosuka City<br />

Museum for information on <strong>the</strong> distribution <strong>of</strong> Recent species <strong>of</strong> <strong>the</strong> genus <strong>Rhinogobius</strong>. I<br />

am also grateful to Dr. Ryuzo Toriyama and Dr. Masamichi Ota <strong>of</strong> <strong>the</strong> Kitakyushu<br />

Museum <strong>of</strong> Natural History for <strong>the</strong>ir constant encouragement.


112 Yoshitaka Yabumoto<br />

Material:<br />

Class Osteichthyes<br />

Order Perciformes<br />

Family <strong>Gobiid</strong>ae<br />

<strong>Rhinogobius</strong> giurinus (Rutter)<br />

Specimens were collected by Akinori Takayama, Teruya Uyeno, and<br />

Yoshitaka Yabumoto between 1983-1986.<br />

KMNH (Kitakyushu Museum <strong>of</strong> Natural History) VP 100,117, almost complete<br />

specimen with dorsal side exposed, but first dorsal fin and anal fin missing. Head region<br />

well preserved. Standard length 19.8 mm and number <strong>of</strong> vertebrae 10+16=26. This<br />

specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> pelvic fin. KMNH VP 100,118, almost complete<br />

specimen with its left side exposed, but second dorsal fin missing. Scales <strong>of</strong> <strong>the</strong> posterior<br />

part <strong>of</strong> <strong>the</strong> body are well preserved. Standard length 24.8 mm and number <strong>of</strong> vertebrae<br />

10+16=26. This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,119,<br />

almost complete specimen with ventral side exposed, but first dorsal fin, a part <strong>of</strong> 2nd<br />

dorsal fin and pelvic fin missing. Standard length 20.6 mm and number <strong>of</strong> vertebrae 10<br />

+ 16=26. This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,120,<br />

almost complete specimen with its right side exposed, but lacking dorsal fin, a part <strong>of</strong> anal<br />

fin and some bones <strong>of</strong> head region. Standard length 26.1 mm and number <strong>of</strong> vertebrae<br />

10+16=26. This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> pelvic fin. KMNH VP<br />

100,121, almost complete specimen with its right side exposed, but lacking pectoral fin,<br />

pelvic fin, anal fin and a part <strong>of</strong> head region. Scales are preserved on abdominal part<br />

and are detached from caudal part. Standard length 25.6 mm. This specimen is<br />

identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,122, almost complete specimen with<br />

its right side exposed. Scales are preserved. Standard length 26.0 mm and number <strong>of</strong><br />

vertebrae 10+16=26. This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH<br />

VP 100,123, anterior part <strong>of</strong> <strong>the</strong> body and caudal fin with its left side exposed, scales<br />

preserved on <strong>the</strong> part below second dorsal fin. Standard length 24.1 mm. This<br />

specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. Takayama's specimen 1, almost<br />

complete specimen with its left side exposed, but lacking a part <strong>of</strong> head region and anal<br />

fin. Scales are preserved. Standard length 27.0 mm and number <strong>of</strong> vertebrae 10+16=<br />

26. This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,124, some<br />

bones <strong>of</strong>head region, pectoral fin, dorsalspines, and scales are preserved. This specimen<br />

is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales.<br />

Distinguishing characters: The branching point <strong>of</strong> <strong>the</strong> innermost pelvic fin ray is<br />

approximately in between <strong>the</strong> base and <strong>the</strong> branching point <strong>of</strong> <strong>the</strong> next ray. The<br />

posterior end <strong>of</strong> each scale is slightly pointed. The spines <strong>of</strong> each scale are short at <strong>the</strong><br />

posterior corner and gradually become longer toward <strong>the</strong> dorsal and ventral ends <strong>of</strong> <strong>the</strong><br />

posterior margin. Grooves extend close to <strong>the</strong> posterior margin <strong>of</strong> <strong>the</strong> scales.<br />

Description: Scales are ctenoid. The frontal bone is wide at <strong>the</strong> posterior part<br />

and rapidlynarrows at <strong>the</strong>anteriorpart. The sensory canalis presenton <strong>the</strong> lateral margin


<strong>Pleistocene</strong> gobiid fishes from Oita 113<br />

preo<br />

max<br />

1mm<br />

Fig. 1. Dorsal view <strong>of</strong> head region <strong>of</strong> a <strong>Pleistocene</strong> <strong>Rhinogobius</strong> giurinus, KMNH VP 100,117.<br />

den, dentary; ecp, ectopterygoid; eth, ethmoid; exo, exoccipital; fro, frontal; max,<br />

maxillary;ope, opercle;pal, palatine; prem, premaxillary; preo, preopercle; pto, pterotic;<br />

qua, quadrate; supo, supraoccipital; sym, symplectic.<br />

<strong>of</strong> <strong>the</strong> frontal and opens at <strong>the</strong> lateral end. The melanin <strong>of</strong> <strong>the</strong> eye ball is preserved in<br />

KMNH VP 100,117 (Fig. 1). The dentary is narrow at <strong>the</strong> anterior part and deep at <strong>the</strong><br />

posterior part. Small unicuspid teeth are present on <strong>the</strong> premaxillary and <strong>the</strong> dentary<br />

(Fig. 1). The ceratohyal is narrow at <strong>the</strong> anterior part and wide at <strong>the</strong> posterior part.<br />

Four branchiostegals are attached on <strong>the</strong> ceratohyal. The epihyal is almost triangular in<br />

shape. The posterior margin <strong>of</strong> <strong>the</strong> pectoral fin is circular. O<strong>the</strong>r observable bones <strong>of</strong><br />

<strong>the</strong> head region are maxillary, quadrate, symplectic, palatine, ectopterygoid, opercle,<br />

preopercle, supraoccipital, pterotic and exoccipital (Fig. 1). The first and second and <strong>the</strong><br />

third and fourth hypurals are fused. The parhypural is short. The haemal spine <strong>of</strong> <strong>the</strong><br />

second preuralcentrum is thick and wide. O<strong>the</strong>r observable bones <strong>of</strong> <strong>the</strong> caudal region<br />

are a part <strong>of</strong> <strong>the</strong> 5th hypural and a part <strong>of</strong> <strong>the</strong> epural (Fig. 2).


114 Yoshitaka Yabumoto<br />

Fig. 2. Caudal region <strong>of</strong> a <strong>Pleistocene</strong> <strong>Rhinogobius</strong> giurinus, KMNH VP 100,122. epu, epural;<br />

hes, haemal spine <strong>of</strong> <strong>the</strong> second preuralcentrum; hyu, hypural; parh, parhypural.<br />

<strong>Rhinogobius</strong> brunneus (Temminck et Schlegel)<br />

Material: KMNH VP 100,125, almost complete specimen with its left side exposed.<br />

Standard length 26.0mm and number <strong>of</strong> vertebrae 10+16=26. This specimen is<br />

identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> pelvic fin. KMNH VP 100,126, almost complete specimen<br />

with its right side exposed. Standard length 24.3 mm and number <strong>of</strong>vertebrae 10+16=<br />

26. This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,127, body<br />

twisted at anal origin, all elements <strong>of</strong> skeleton, fin rays and scales are detached. This<br />

specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,128, almost complete<br />

specimen with its left side exposed, but ribs, pectoral, pelvic and anal fins missing.<br />

Standard length 23.7 mm. This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales.<br />

KMNH VP 100,129, almost complete specimen with its left side exposed, but pelvic fins<br />

missing. Standard length 35.8 mm and number <strong>of</strong> vertebrae 10+16=26. This speci<br />

men is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,130, specimen with its left


<strong>Pleistocene</strong> gobiid fishes from Oita 115<br />

side exposed lacking first dorsal fin, caudal skeleton and caudal fin. This specimen is<br />

identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,131, specimen with ventral side<br />

exposed lacking caudal part. Some bones <strong>of</strong> head region preserved. This specimen is<br />

identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. Takayama's specimen 2, almost complete specimen<br />

with its left side exposed, but first dorsal fin missing. Some bones <strong>of</strong> head region<br />

disarticulated. Standard length 37.5 mm and number <strong>of</strong> vertebrae 10+16=26. This<br />

specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,132, almost complete<br />

specimen with its right side exposed, but a part <strong>of</strong> <strong>the</strong> anal fin missing. Standard length<br />

37.4 mm and number <strong>of</strong> vertebrae 10+16=26. This specimen is identified by <strong>the</strong> form<br />

<strong>of</strong> <strong>the</strong> scales. KMNH VP 100,133, almost complete specimen with its left side exposed,<br />

and some bones <strong>of</strong> head region disarticulated. Standard length 27.0 mm and number <strong>of</strong><br />

vertebrae 10+16=26. This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH<br />

VP 100,134, almost complete specimen with its left side exposed, but lacking some bones<br />

<strong>of</strong> head region and a part <strong>of</strong> abdominal vertebrae. This specimen is identified by <strong>the</strong><br />

form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,135, caudal part <strong>of</strong> <strong>the</strong> body. This specimen is<br />

identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,136, anterior part <strong>of</strong> <strong>the</strong> body.<br />

This specimen is identified by <strong>the</strong> form <strong>of</strong> <strong>the</strong> scales. KMNH VP 100,137, caudal part <strong>of</strong><br />

<strong>the</strong> body, lacking upper part <strong>of</strong> <strong>the</strong> caudal skeleton.<br />

This specimen is identified by <strong>the</strong><br />

form <strong>of</strong> <strong>the</strong> scales.<br />

Distinguishing characters: The branching point <strong>of</strong> <strong>the</strong> innermost pelvic fin ray is<br />

beside <strong>the</strong> branching point <strong>of</strong> <strong>the</strong> next ray. The posterior margin <strong>of</strong> each scale is round<br />

in shape. The spines <strong>of</strong> each scale are almost equal in length and grooves extend two<br />

pro<br />

prem<br />

den<br />

max<br />

Fig.3. Ventral view <strong>of</strong> head region <strong>of</strong> a <strong>Pleistocene</strong> <strong>Rhinogobius</strong> brunneus, KMNH VP<br />

100,131. ang, angular; bra, branchiostegal; para, parashenoid; pro, prootic; o<strong>the</strong>r<br />

abbreviations see Fig. 1.


116 Yoshitaka Yabumoto<br />

thirds <strong>of</strong> <strong>the</strong> way from <strong>the</strong> anterior end.<br />

Description: Scales are ctenoid. In <strong>the</strong> fossil specimens, <strong>the</strong> observable bones <strong>of</strong><br />

<strong>the</strong> head region are maxillary, premaxillary, dentary, angular, palatine, ectopterygoid,<br />

quadrate, symplectic, frontal, parasphenoid, prootic, and branchiostegals. There are two<br />

condyles on <strong>the</strong> anterior end <strong>of</strong> <strong>the</strong> maxillary. The middle part <strong>of</strong> <strong>the</strong> maxillary is broad.<br />

Small unicuspid teeth are present on <strong>the</strong> premaxillary and <strong>the</strong> dentary. The palatine has<br />

two processes and one acetabulum at <strong>the</strong> anterior end. The posterior margin connects<br />

with <strong>the</strong> ectopterygoid which is pointed at <strong>the</strong> upper end. The lower end <strong>of</strong> <strong>the</strong><br />

ectopterygoid is wide and connects with <strong>the</strong> anterior margin <strong>of</strong> <strong>the</strong> quadrate which is<br />

approximately triangular with a long process ventrally. The antero-ventral corner <strong>of</strong> <strong>the</strong><br />

quadrate is a condyle for <strong>the</strong> angular (Fig. 3). A pair <strong>of</strong> foramina is present at <strong>the</strong> middle<br />

<strong>of</strong> <strong>the</strong> parasphenoid. The anterior quarter <strong>of</strong> <strong>the</strong> ventral surface <strong>of</strong> <strong>the</strong> parasphenoid is<br />

slightly concave for <strong>the</strong> attachment to <strong>the</strong> prevomer. The prootichas a large foramen for<br />

<strong>the</strong> facial nerve (Fig. 3). Forms <strong>of</strong> <strong>the</strong>se observable bones are similar to those <strong>of</strong> Recent<br />

R. brunneus.<br />

Remarks: In <strong>the</strong> Recent species <strong>of</strong> <strong>the</strong> genus <strong>Rhinogobius</strong>, <strong>the</strong> first pair <strong>of</strong> normal<br />

long ribs are attached to <strong>the</strong> third vertebra. The number <strong>of</strong> dorsal pterygiophores<br />

correspond to <strong>the</strong> number<strong>of</strong>dorsal fin rays. The space between <strong>the</strong> fifth and sixthspines<br />

<strong>of</strong> <strong>the</strong> first dorsal fin is much wider than spaces between o<strong>the</strong>r spines <strong>of</strong> <strong>the</strong> fin. The<br />

number <strong>of</strong> anal pterygiophores is one fewer than that <strong>of</strong>anal fin rays. The haemalspine<br />

<strong>of</strong> <strong>the</strong> first caudal vertebra is conspicuous (Fig.6). This information is useful in<br />

determining <strong>the</strong> numbers <strong>of</strong> abdominal and caudal vertebrae, and <strong>the</strong> numbers <strong>of</strong> dorsal<br />

and anal fin rays in <strong>Rhinogobius</strong> fossil material.<br />

The following characters indicate that <strong>the</strong> fossil specimens are members <strong>of</strong> <strong>the</strong><br />

teleostean fish family <strong>Gobiid</strong>ae <strong>of</strong> <strong>the</strong> order Perciformes: (1) <strong>the</strong> first and second dorsal<br />

fins are present and are notcontinuous; (2) both pelvic fins are close toge<strong>the</strong>r andsituated<br />

below <strong>the</strong> pectoral fin; (3) <strong>the</strong> body is covered with ctenoid scale; (4) <strong>the</strong> hypural bones<br />

arefused and simplified; (5) <strong>the</strong> dentary and premaxillary bear small conical teeth; (6) <strong>the</strong><br />

interorbital space is narrow. They have <strong>the</strong> following meristic characters: (1) <strong>the</strong><br />

number <strong>of</strong> abdominal vertebrae is 10 and <strong>the</strong> number <strong>of</strong> caudal vertebrae is 16; (2) <strong>the</strong><br />

first dorsal fin consists <strong>of</strong>6 spines and <strong>the</strong>second dorsal fin consists <strong>of</strong>onespine and 8 s<strong>of</strong>t<br />

rays; (3) <strong>the</strong> anal fin also consists <strong>of</strong>onespine and 8 s<strong>of</strong>t rays. According to Masuda et<br />

al. (1984), 35 species in 18 genera <strong>of</strong> gobiid fishes are found in freshwater in Japan.<br />

Among <strong>the</strong>m, <strong>the</strong> meristic characters <strong>of</strong> <strong>Rhinogobius</strong> giurinus and R. brunneus correspond to<br />

that <strong>of</strong> <strong>the</strong> fossil specimens. Threespecies <strong>of</strong><strong>the</strong> genus <strong>Rhinogobius</strong> have been reported in<br />

Japan. The fossil specimens are not <strong>Rhinogobius</strong> flumineus, because R. fiumineus has 11<br />

abdominal vertebrae. The differences between Recent R. giurinus and R. brunneus are<br />

recognized in <strong>the</strong>forms <strong>of</strong><strong>the</strong>sucking disk <strong>of</strong><strong>the</strong> pelvic fin andscales. Uyeno and Iwao<br />

(1966) described <strong>the</strong> differences in <strong>the</strong> shape <strong>of</strong> <strong>the</strong> sucking disk and <strong>the</strong> number <strong>of</strong><br />

branches <strong>of</strong> <strong>the</strong> innermost pelvic fin ray. But <strong>the</strong>se characters <strong>of</strong> <strong>the</strong> pelvic fin are not


<strong>Pleistocene</strong> gobiid fishes from Oita 117<br />

1mm<br />

CSs"^<br />

1mm<br />

Fig. 4. Pelvic fins <strong>of</strong> Recent and fossil specimens <strong>of</strong> <strong>the</strong> genus <strong>Rhinogobius</strong>. A, Recent R.<br />

giurinus; B, Recent R. brunneus; C, fossil R. giurinus, KMNH VP 100,118; D, fossil R.<br />

brunneus, KMNH VP 100,125. The lines indicate <strong>the</strong> branching points <strong>of</strong> fin rays.<br />

1mm<br />

Fig. 5. Scales <strong>of</strong> Recent and fossil specimens <strong>of</strong> <strong>the</strong> genus <strong>Rhinogobius</strong>. A and B, Recent R.<br />

giurinus; C and D, Recent R. brunneus; E, fossil R. giurinus, KMNH VP 100,118; F,<br />

fossil R. brunneus, KMNH VP 100,132.


<strong>Pleistocene</strong> gobiid fishes from Oita 119<br />

observable in <strong>the</strong> fossil specimens from Kusu Basin, because <strong>the</strong> posterior part <strong>of</strong> <strong>the</strong><br />

pelvic fin is not preserved. In Recent R. giurinus and R. brunneus, <strong>the</strong>re is a difference in<br />

<strong>the</strong> position <strong>of</strong> <strong>the</strong> branching point <strong>of</strong> <strong>the</strong> innermost ray <strong>of</strong> <strong>the</strong> pelvic fin. The branching<br />

point <strong>of</strong> <strong>the</strong> innermost ray is approximately in between <strong>the</strong> base and <strong>the</strong> branching point<br />

<strong>of</strong> <strong>the</strong> next ray in R. giurinus (Fig. 4A). The branching point <strong>of</strong> <strong>the</strong> innermost ray is<br />

beside to <strong>the</strong> branching point <strong>of</strong> <strong>the</strong> next ray in R. brunneus (Fig. 4B). This character is<br />

observable in <strong>the</strong> fossil specimens (Fig. 4C and D).<br />

In R.giurinus, <strong>the</strong> posterior end <strong>of</strong> <strong>the</strong> scale is slightly pointed and <strong>the</strong> spines are short<br />

on <strong>the</strong> posterior corner and gradually become longer toward <strong>the</strong> dorsal and ventral ends <strong>of</strong><br />

<strong>the</strong> posteriormargin. Grooves extendclose to <strong>the</strong> posteriormargin <strong>of</strong> <strong>the</strong> scale (Fig.5B).<br />

In R. brunneus, <strong>the</strong> posterior margin <strong>of</strong> <strong>the</strong> scale is round in shape and <strong>the</strong> spines are<br />

almostequal in length. Grooves extend two thirds<strong>of</strong> <strong>the</strong> way from <strong>the</strong> anterior end (Fig.<br />

5C). In R. giurinus, scales similar to those <strong>of</strong> R. brunneus occur in a ratio <strong>of</strong> approximately<br />

one to twenty (Fig. 5A). In R. brunneus, scales similar to those <strong>of</strong> R. giurinus occur in a<br />

ratio <strong>of</strong> one to five or six (Fig. 5D). Two types <strong>of</strong> scales are recognized in <strong>the</strong> fossil<br />

specimens (Fig. 5E and F). All <strong>of</strong> <strong>the</strong> observablescales in a fossil specimen are examined<br />

for identification <strong>of</strong> species. Two species, R. giurinus and R. brunneus, are recognized<br />

among fossil specimens on <strong>the</strong> bases <strong>of</strong> <strong>the</strong> characters <strong>of</strong> <strong>the</strong> pelvic fin and scales (Figs. 4C<br />

and D, 5E and F).<br />

Literature Cited<br />

Iwauchi, A. and Y. Hase. 1987. Late Cenozoic vegetation and paleoenvironment <strong>of</strong> nor<strong>the</strong>rn and<br />

central Kyushu, Japan. Part 3. Sou<strong>the</strong>rn part <strong>of</strong> Kusu Basin (Lower and Middle <strong>Pleistocene</strong>).<br />

Jour. Geol. Soc. Japan, 93 (7): 469-489, figs. 1-13, tabs. 1-2, pis. 1-2.<br />

Masuda, H., K. Amaoka, C. Araoa, T. Uyeno, and T. Yoshino (ed.). 1984. The fishes <strong>of</strong> <strong>the</strong><br />

Japanese Archipelago. Text. Tokai Univ. Press, xxii+437 pp., 245 figs.<br />

Shuto, T. 1953. Younger Cenozoic history <strong>of</strong> Oita District, Kyushu (I). Jour. Geol. Soc. Japan, 59<br />

(693): 225-240, figs. 1-8, tabs. 1-6.<br />

Uyeno, T., S. Kimura, and Y. Hasegawa. 1975. Freshwater fishes from Late Cenozoic deposits in<br />

Kusu Basin, Oita Prefecture, Japan. Mem. Natn. Sci. Mus. (8): 57-66, fig. 1, tabs. 1-2, pis. 6-9.<br />

Uyeno, T. and Y. Iwao. 1975. Late Cenozoic gobiid fish from Togo Formation in Kagoshima<br />

Prefecture, Japan. Bull. Natn. Sci. Mus., Ser. C (Geol.), 1 (2): 55-60, figs. 1-2, pis. 1-2.


<strong>Pleistocene</strong> <strong>Gobiid</strong> <strong>Fishes</strong> <strong>of</strong> <strong>the</strong> <strong>Genus</strong> <strong>Rhinogobius</strong><br />

from Kusu Basin, Oita Prefecture, Japan.<br />

Yoshitaka Yabumoto<br />

Plates 1-2


Explanation <strong>of</strong> Plate 1.<br />

Photographs <strong>of</strong> <strong>the</strong> fossils <strong>of</strong> <strong>Rhinogobius</strong> giurinus (Rutter) from Kusu<br />

Basin, Oita Prefecture.<br />

A. lateral view <strong>of</strong> <strong>the</strong> left side (KMNH VP 100,118).<br />

B. ventral view (KMNH VP 100,119).<br />

C dorsal view (KMNH VP 100,117).<br />

Scales indicate 10 mm.


Yabumoto, Y. <strong>Pleistocene</strong> gobiid fishes from Oita Plate 1<br />

wtmm...<br />

B<br />

C


Plate2<br />

I ;•••; !


Explanation <strong>of</strong> Plate 2.<br />

Photographs <strong>of</strong> <strong>the</strong> fossils <strong>of</strong> <strong>Rhinogobius</strong> brunneus (Temminck et Schlegel)<br />

from Kusu Basin, Oita Prefecture.<br />

A. lateral view <strong>of</strong> <strong>the</strong> left side (KMNH VP 100,125).<br />

B. lateral view <strong>of</strong> <strong>the</strong> left side (KMNH VP 100,129).<br />

C lateral view <strong>of</strong> <strong>the</strong> right side (KMNH VP 100,132).<br />

Scales indicate 10 mm.


Yabumoto, Y. <strong>Pleistocene</strong> gobiid fishes from Oita Plate 2<br />

B<br />

irp

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